Distribution of Lithocysts, Trichomes, Hydathodes and Stomata in Leaves of Pilea cadierei Gagnep. & Guill. (Urticaceae)

In leaves of Pilea cadierei lithocysts, trichomes and hydathodesin the upper epidermis, and lithocysts, trichomes and anomocyticstomata in the lower epidermis are components of a regular distributionpattern in which spacing probably depends upon inhibitory interactionsbetween initial cells during leaf development. Anisocytic stomata,which also occur in the lower epidermis, show a clumped distributionpattern that results from their relatively late formation aftercell division in the epidermis is otherwise complete, and fromthe inhibitory influence of established lithocysts, trichomesand anomocytic stomata. Ordered spacing of anisocytic stomatawithin groups probably results from mutual inhibition betweeninitial cells at the time of inception. Lithocysts associatedwith primary and secondary veins are elongated along the veinaxis. Elsewhere in the leaf lithocyst, orientation is obliquerelative to the midrib and shows a correlation with vein orientationaround the time of lithocysts elongation. Lithocyst, cystolith, trichome, hydathode, stomata, Pilea cadierei     

Release of Guttation Fluid from Passive Hydathodes of Intact Barley Plants. I. Structural and Cytological Aspects

The structural details of the guttating tips of 7-day-old barleyleaves were studied as a basis for a subsequent report on thephysiology of guttation. The walls of the vessels at the tipsof leaves bear many pits and are rather thin, appearing neithercutinized nor lignified. This could facilitate a direct passageof solutes out of the xylary system through the leaf apoplastand out to the leaf surface via hydathode openings. The latterare formed by modified stomatal guard cells, and there are nospecially differentiated epithem, epithelium or gland hair likestructures that could serve an active elimination of guttation.Xylem parenchyma cells and the peculiar mesophyll cells withdense cytoplasm, numerous mitochondria, an extended ER systemand a considerable formation of small vesicles in the leaf tipcould modify the content of the guttated fluid along the routeof transport. Hordeum vulgare L., barley, hydathodes, guttation     

Hydathode anatomy in Potentilla palustris (Rosaceae)

Guttating leaf teeth of Potentilla palustris plants from Wisconsin, USA, were cleared or processed for plastic sectioning or scanning electron microscopy. Anatomical features include: 1) long slender hydathode area occupying most of the tooth, 2) adaxial pad of small, flat epidermal cells with 50 or more sunken water pores about the size of ordinary abaxial stomates, 3) three converged bundles that extend distally, where their tracheary files are separated by intervening files of xylem parenchyma cells with sinuous walls, 4) adaxial mass of small, loosely arranged epithem cells above the xylem, 5) one slender phloem strand that extends only about a third of the way into the hydathode, and 6) bundle sheath extending distally only abaxially and along the flanks of the hydathode. Potentilla hydathodes differ significantly from non-guttating ones described earlier in Physocarpus (Rosaceae).     

Ultrastructure of hydathode trichomes of hemiparasitic Rhinanthus alectorolophus and Odontites vernus: how important is their role in physiology and evolution of parasitism in Orobanchaceae?

The Rhinanthoid clade of the family Orobanchaceae comprises plants displaying a hemiparasitic or holoparasitic strategy of resource acquisition. Some of its species (mainly Rhinanthus spp.) are often used as models for studies of hemiparasite physiology. Although there is a well‐developed concept covering their physiological processes, most recent studies have neglected the existence of hydathode trichomes present on leaves of these hemiparasitic plants. As a first step for the proposed integration of these structures in the theory of physiological processes of the hemiparasites, we described the outer micromorphology and ultrastructure of the hydathode trichomes on leaves of hemiparasitic Rhinanthus alectorolophus and Odontites vernus with scanning and transmission electron microscopy (SEM and TEM, respectively). The TEM inspections of both types of trichome revealed typical ultrastructural features: labyrinthine cell wall, high content of cytoplasm in cells with numerous mitochondria and presence of plasmodesmata. All these features indicate high metabolic activity complying with their function as glandular trichomes actively secreting water. The active secretion of water by the hydathode trichomes (evidence for which is summarised here) also presents a possible mechanism explaining results of previous gas exchange measurements detecting high dark respiration and transpiration rates and a tight inter‐correlation between them in hemiparasitic Orobanchaceae. In addition, this process is hypothesised to have allowed multiple evolutionary transitions from facultative to obligate hemiparasitism and unique xylem‐feeding holoparasitism of Lathraea with a long‐lived underground stage featuring a rhizome covered by scales of leaf origin.     

Hydathode trichomes actively secreting water from leaves play a key role in the physiology and evolution of root-parasitic rhinanthoid Orobanchaceae

Background and Aims Root hemiparasites from the rhinanthoid clade of Orobanchaceae possess metabolically active glandular trichomes that have been suggested to function as hydathode trichomes actively secreting water, a process that may facilitate resource acquisition from the host plant’s root xylem. However, no direct evidence relating the trichomes to water secretion exists, and carbon budgets associated with this energy-demanding process have not been determined.Methods Macro- and microscopic observations of the leaves of hemiparasitic Rhinanthus alectorolophus were conducted and night-time gas exchange was measured. Correlations were examined among the intensity of guttation, respiration and transpiration, and analysis of these correlations allowed the carbon budget of the trichome activity to be quantified. We examined the intensity of guttation, respiration and transpiration, correlations among which indicate active water secretion.Key Results Guttation was observed on the leaves of 50 % of the young, non-flowering plants that were examined, and microscopic observations revealed water secretion from the glandular trichomes present on the abaxial leaf side. Night-time rates of respiration and transpiration and the presence of guttation drops were positively correlated, which is a clear indicator of hydathode trichome activity. Subsequent physiological measurements on older, flowering plants indicated neither intense guttation nor the presence of correlations, which suggests that the peak activity of hydathodes is in the juvenile stage.Conclusions This study provides the first unequivocal evidence for the physiological role of the hydathode trichomes in active water secretion in the rhinanthoid Orobanchaceae. Depending on the concentration of organic elements calculated to be in the host xylem sap, the direct effect of water secretion on carbon balance ranges from close to neutral to positive. However, it is likely to be positive in the xylem-only feeding holoparasites of the genus Lathraea, which is closely related to Rhinanthus. Thus, water secretion by the hydathodes might be viewed as a physiological pre-adaptation in the evolution of holoparasitism in the rhinanthoid lineage of Orobanchaceae.     

Laminar hydathodes inUrticaceae: Survey of tribes and anatomical observations onPilea pumila andUrtica dioica

Laminar hydathodes are known from only three dicot families. InUrticaceae they are associated with minor vein junctions in all five tribes, as surveyed from cleared leaves of 43 species in 30 genera. Only one species lacked hydathodes. Exclusively adaxial hydathodes were found in 28 genera. In tribeElatostemeae, laminar hydathodes inPilea andPellionia species are abaxial, adaxial, or on both surfaces. Guttation was observed in four species.Urtica dioica (adaxial) andPilea pumila (abaxial) were studied anatomically in detail. Hydathodes in the former have normal bundle structure but xylem gaps sometimes occur. In the latter, phloem is displaced in three previously undescribed ways: 1) ends abruptly near hydathode, 2) curves into connecting vein at adjacent junction, or 3) departs xylem, skirts hydathode independently, and rejoins adjacent xylem strand. Laminar hydathodes are a unifying character of theUrticaceae, and they also strengthen its close relationship to theMoraceae.     

ONTOGENY OF THE TENDRIL IN VITIS VINIFERA

The apical meristem of the grape tendril possesses several remarkable features: bilateral symmetry, a minimal number of appendages, and an exceedingly brief period of apical meristem activity. The cellular configuration of the apex changes from tunica-corpus to zonate, as rudimentary leaves and branch-tendril apices are initiated. Eventually the apical meristem of the tendril itself ceases meristematic activity and differentiates as a large hydathode. Typical spongy epithem tissue, copious xylary tissue, and water-pores in the epidermis characterize the hydathode. Numerous vascular strands traverse the length of the tendril and terminate in enlarged tracheary elements adjacent to the epithem. Cessation of meristematic activity follows lowered mitotic rate in the summit and accelerated differentiation below and within the meristem.     

Ion gradients in xylem exudate and guttation fluid related to tissue ion levels along primary leaves of barley

The concentration of ions in plant cells and tissues is an essential factor in determining physiological function. In the present study, we established that concentration gradients of mobile ions exist in both xylem exudates and tissues within a barley (Hordeum vulgare) primary leaf. For K⁺ and NO₃^?, ion concentrations generally decreased from the leaf base to the tip in both xylem exudates and tissues. Ion gradients were also found for Pi and Cl^? in the xylem. The hydathode strongly absorbed Pi and re‐translocated it to the rest of the plant, whereas Cl^? was extruded. The ion concentration gradients developed early during leaf growth, increased as the tissue aged and remained under both high and low transpiration conditions. Measurement of the expression profiles of Pi, K⁺ and NO₃^? transporters along the longitudinal axis of the leaf revealed that some transporters are more expressed at the hydathode, but for most transporters, there was no significant variation along the leaf. The mechanisms by which longitudinal ion gradients develop in leaves and their physiological functions are discussed.     

Marginal and laminar hydathode-like structures in the leaves of the desiccation-tolerant angiosperm Myrothamnus flabellifolius Welw.

The fan-shaped leaves of the resurrection plant Myrothamnus flabellifolius Welw. fold during episodes of drought and consequent desiccation of the tissue. The leaf teeth of M. flabellifolius have several features characteristic of hydathodes. Tracheary elements from the three vein endings that converge in each leaf tooth subtend and extend into a cluster of cells significantly smaller than those of the adjacent mesophyll. The stomata overlying this putative epithem are larger than the other stomata on the leaf surface. Crystal violet is absorbed via these stomata in non-transpiring leaves, suggesting that they are water pores. Two to four such water pores occur per hydathode and are readily distinguished in desiccated leaves. Laminar hydathodes apparently also occur in the leaves of M. flabellifolius. Branched vein endings that terminate in short, wide tracheary elements subtend the outer edges of the abaxial leaf ridge, which otherwise lack stomata, and coincide with regions of crystal violet uptake. Guttation could not be induced in M. flabellifolius. However, desiccated leaves readily absorb liquid water through the leaf surface. The use of Calcafluor White to trace the pathway of apoplastic water movement suggests a role for both types of hydathode in foliar water uptake during rehydration while the accumulation of Sulphorhodamine G (indicating solute retrieval from the apoplast) in the epithem of transpiring plants suggests the hydathodes may be a pathway of water loss in the desiccating leaf.     

The bHLH protein, MUTE, controls differentiation of stomata and the hydathode pore in Arabidopsis

Stomata are turgor-driven epidermal valves on the surface of plants that allow for efficient gas and water exchange between the plant and its environment. The Arabidopsis thaliana basic helix-loop-helix (bHLH) protein, MUTE, is a master regulator of stomatal differentiation where it is required for progression through the stomatal lineage and the differentiation of stomata. The genetic control of stomatal spacing across the epidermal surface is variable in different organs. For instance, a distinct suite of genes from those in leaves regulates stomatal patterning in hypocotyls. Here we report that regardless of organ type, MUTE controls downstream events directing stomatal differentiation, specifically the transition from meristemoid to guard mother cell. Ectopic MUTE expression is sufficient to over-ride cell fate specification in cell types that do not normally differentiate stomata. Furthermore, MUTE is required for the production of the structure evolutionarily related to stomata, the hydathode pore. Consistently, MUTE displays expression at the tip of cotyledons and leaves, thus co-localizing with the auxin maxima. However, MUTE itself was not regulated by the auxin, and the absence of hydathode pores in mute did not affect the auxin maxima. Surprisingly, our analysis revealed that the requirement for MUTE could be partially circumvented under conditions of compromised inhibitory signaling.     

Electrophoretic Variation in Esterases, Peroxidases and Acid Phosphatases in Some Native Greek Taxa of the Genera Hordeum and Taeniatherum

Horizontal starch gel electrophoresis has been applied to thestudy of esterase, peroxidase and acid phosphatase patternsin seven taxa, namely Hordeum diploids (2n=14) (H. marinum,H. marinum I and H. hystrix), tetraploids (2n=28) (H. bulbosumand H. murinum subsp. leporinum) and Taeniatherum (2n=14) (T.caput-medusae and T. caput-medusae I) in order to elucidatetheir phylogenetic relationships. On the basis of our experimentalresults the seven taxa may be placed in the following threegroups; (1) diploid Hordeum (H. marinum, H. marinum I, H. hystrix);(2) tetraploid Hordeum (H. bulbosum, H. murinum subsp. leporinum);(3) Taeniatherum (T. caput-medusae, T. caput-medusae I). Esterase, peroxidase and acid phosphatase patterns of the twoHordeum diploid taxa (H. marinum and H. marinum I) are verysimilar suggesting their close phylogenetic relationship; thesame is true for both the taxa of the genus Taeniatherum (T.caput-medusae and T. caput-medusae I). The taxa of the Taeniatherumgroup compared with the diploid Hordeum (H. marinum, H. marinumI, H. hystrix) and the tetraploid Hordeum (H. bulbosum, H. murinumsubsp. leporinum) show a lower degree of phylogenetic relationshipand seem to be equally distant from them. The tetraploid Hordeumgroup shows a higher phylogenetic relationship with diploidHordeum group than with the Taeniatherum group. These results confirm that the genus Taeniatherum, previouslyconsidered as part of the genus Hordeum, should be regardedas a separate genus. Gramineae (Poaceae), Hordeum L., Taeniatherum Nevski., esterase, peroxidase and acid phosphatase patterns, phylogenetic relationships     

The Regulatory Functions of the rolB and rolC Genes of Agrobacterium rhizogenes are Conserved in the Homologous Genes (Ngrol) of Nicotiana glauca in Tobacco Genetic Tumors

To compare patterns of expression between the Ngrol genes ofN. glauca and the Rirol genes of Agrobacterium rhizogenes, weperformed fluorometric and histochemical analysis of transgenicgenetic tumors on the hybrid of Nicotiana glauca x N. langsdorffü(Fl) that harbored a rß- glucuronidase (GUS) reportergene fused to the promoter of NgrolB, NgrolC, RirolB or RirolC The promoters of NgrolB and NgrolCNgrolC had 2- to 3-fold loweractivity than those of RirolB and RirolC However, the changesin patterns of GUS activity caused by deletion of NgrolB andNgrolCpromoters were similar to those of RirolB and RirolC promoters.This result suggests that the cis-acting sequences that regulatethe level of expression of RirolB and RirolC are conserved inthe NgrolB and NgrolC promoters. Furthermore, an auxin dependent(NAA-dependent) increase in GUS activity was observed in thecase of NgrolB-GUS and RirolB-GUS. Histochemical analysis showedGUS activity encoded by both NgrolB-GUS and RirolB-GUS in normal-typeFl transgenic plants was located in meristematic zones, whilethat encoded by NgrolC-GUS and RirolC-GUS was detected mainlyin vascular systems of various organs. Thus, the patterns ofexpression of the Ngrol genes were the same as those of theRirol genes in terms of promotion by auxin and tissue-specificity,indicating that regulatory mechanisms for both sets of geneshave been conserved during the evolution of the genus Nicotianaafter transfer from a progenitor of Agrobacterium to that ofNicotiana. (Received May 2, 1995; Accepted June 13, 1995)     

MORE MOLLUSCAN NAME-CHANGES, GENERIC AND SPECIFIC

Volema, Bolten, type V. paradisiaca, Bolten, has priority overMelongena, Schumacher, but may be used independently. Mayena, gen. nov., proposed for Biplex australasia, Perry. Turricula, Schumacher, is the correct name for Surcula, H. &A. Adams. Gelagna, Schaufuss, is equal to and antedates Paralagena, Dall. Partulida, Schaufuss, should replace Spiralinella, Chaster. Campanile, Fischer, has for type the recent species Cerithiumleve, Quoy & Gaimard, which is here renamed Campanile symbolicum,sp. nov. Campanilopa, gen. nov., introduced for the fossil Cerithiumgiganteum. Lam. Pleurotomoides, Bronn, has priority over Lora, Gistel, and Clathurella,Carpenter, all proposed as alternatives for Defrancia, Millet,preoccupied. Gabrielona, gen. nov., proposed for Phasianella nepeanensis,Gatliff & Gabriel. Orbitestella, gen. nov., for Cyclostrema bastowi, Gatliff. Megathura, Pilsbry, should be used instead of Macrochasma, Dall. Mitromorpha, auctt., is not Mitromorpha, Carpenter, which wasbased on Daphnella (?) filosa, Carpenter. Antimitra, gen. nov., is proposed for Pleurotoma agrota, Reeve,with which A. Adams' Mitromorpha lirata is congeneric. Lovellona, gen. nov., type Conus atramentosus. Reeve. Apaturris, gen. nov., type Mitramorpha expeditionis, Oliver. Callanaitis, gen. nov., type Venus yatei, Gray, for Salacia,Jukes-Browne, preoccupied. Anopsia, Gistel, is available for Psyche, Rang, preoccupied,and has priority over Verrillopsyche, Cossmann, proposed forHalopsyche, Kieferstein, preoccupied, introduced as substitutefor Rang's name. Hydromyles, Gistel, should be used for Euribia, Rang, preoccupied,as it is older than Kieferstein's name Theceurybia, for thesame genus. (Received 13 April 1917;     

Reproductive Isolation in a Native Population of Petunia sensu Jussieu (Solanaceae)

Reproductive isolation was studied in four syntopic speciesof Petunia sensu Jussieu (Solanaceae) at a site in Rio Grandedo Sul State, Brazil. Reciprocal artificial crossing experimentsconfirmed that a genetic barrier exists between Petunia(P. axillarisand P. integrifolia) andCalibrachoa (C. parviflora and C. heterophylla),and also between C. parviflora andC. heterophylla . Petuniaaxillaris(white, nocturnally scented flower) is geneticallycompatible with the syntopic and allotopic P. integrifolia(coloured,unscented flower). Reproductive isolation appears to be maintainedby the two species having different pollinators: nocturnallyactive hawkmoths (Manduca contracta andM. diffissa subsp. petuniae)pollinate P. axillaris while a diurnally active bee (Hexanthedasp.) pollinates P. integrifolia. Flowers of P. integrifoliaexhibit diurnal opening and closing movements synchronous withthe activity period of the bee. Other than a probable nectarrobber (a carpenter bee, Xylocopa augusti), no insect visitedflowers of P. axillaris in the day. Amounts of floral nectarin P. axillaris and P. integrifolia were within the range ofhawkmoth- and bee-pollinated flowers, respectively. Reproductiveisolating mechanisms in the genus Petunia sensu Jussieu arediscussed. Copyright 2001 Annals of Botany Company Calibrachoa, hawkmoth, Petunia, Petunia axillaris, Petunia integrifolia, pollinator, reproductive isolation, Solanaceae     

Estimating the capacity for improvement in risk prediction with a marker

Consider a set of baseline predictors X to predict a binaryoutcome D and let Y be a novel marker or predictor. This paperis concerned with evaluating the performance of the augmentedrisk model P(D = 1|Y,X) compared with the baseline model P(D= 1|X). The diagnostic likelihood ratio, DLR_X(y), quantifiesthe change in risk obtained with knowledge of Y = y for a subjectwith baseline risk factors X. The notion is commonly used inclinical medicine to quantify the increment in risk predictiondue to Y. It is contrasted here with the notion of covariate-adjustedeffect of Y in the augmented risk model. We also propose methodsfor making inference about DLR_X(y). Case–control studydesigns are accommodated. The methods provide a mechanism toinvestigate if the predictive information in Y varies with baselinecovariates. In addition, we show that when combined with a baselinerisk model and information about the population distributionof Y given X, covariate-specific predictiveness curves can beestimated. These curves are useful to an individual in decidingif ascertainment of Y is likely to be informative or not forhim. We illustrate with data from 2 studies: one is a studyof the performance of hearing screening tests for infants, andthe other concerns the value of serum creatinine in diagnosingrenal artery stenosis.     

Intergeneric and Interspecific Relationships in Araceae tribe Caladieae and Development of Molecular Markers using Amplified Fragment Length Polymorphism (AFLP)

Since amplified fragment length polymorphism (AFLP) analysishas proved useful in distinguishing cultivars of Caladium, itwas used to assess the status of species of Caladium vs. Xanthosoma,both in tribe the Caladieae, and to reassess the position ofHapaline in the same tribe. AFLP analysis using three primercombinations was carried out on four species of Caladium(C.bicolor, C. humboldtii, C. lindenii and C. schomburgkii). Resultsshowed that AFLP can distinguish between the different speciesby their unique and different banding patterns. AFLP analysisconfirmed that C. humboldtii is a species distinct from C. bicolorand that C. lindenii is a true Caladium species and does notbelong to Xanthosoma. UPGMA cluster analysis showed that C.bicolor and C. schomburgkii are most similar and that C. humboldtiiis closer to the C. bicolor / C. schomburgkii cluster comparedwith C. lindenii. Genetic relationships between Caladium, Xanthosoma,Hapaline, Alocasia and Protarum were also examined by AFLP analysisusing eight primer combinations. Several useful molecular markerswere specific either to Caladium orXanthosoma , so that AFLPcan be used to distinguish species of these two genera. Geneticanalysis of the genera examined confirms that the Caladieaeand Colocasieae tribes are distinct and that Hapaline fallswithin the tribe Caladieae and that Protarum is most distantfrom all the genera examined. Copyright 2000 Annals of BotanyCompany Araceae, Caladium, Xanthosoma, Hapaline, Alocasia, Protarum, AFLP DNA fingerprinting, diversity, AFLP markers     

Relationships Between Climate and Flowering of Eight Herbs in a Swedish Deciduous Forest

Effects of annual variation in rainfall, temperature and humidityon flowering abundance of eight temperate woodland plants (Anemonenemorosa, Cardamine bulbifera, Lamiastrum galeobdolon,Oxalisacetosella , Ranunculus ficaria, Stellaria holostea, Viola reichenbachianaand Viola riviniana) were studied during 12 consecutive years(1989–2000) in a hornbeam (Carpinus betulus) forest insoutheast Sweden. Above-average rainfall/humidity in late summerto early autumn of the preceding year increased flowering abundancein L. galeobdolon, O. acetosella, V. reichenbachiana, V. rivinianaand, especially, in R. ficaria, but not in S. holostea and A.nemorosa. Moreover, flowering of R. ficaria and O. acetosellawas positively related to rainfall/humidity during several partsof, or the entire, preceding year. On the contrary, floweringof S. holostea and A. nemorosa was closely related to low valuesof rainfall/humidity in autumn and/or winter of the precedingyear and also to low humidity in the current year in A. nemorosa.Two long periods (3–4 years) of increasing rainfall deficitcoincided with decreasing flowering abundance in most of thespecies, but not with decreasing vegetative development. Temperaturevariability was less consistently related to flowering. A coolperiod during the preceding summer or autumn seemed importantfor flowering in L. galeobdolon, O. acetosella and the Violaspecies, although these relations were, at least partly, causedby interactions with rainfall/humidity. No significant (P <0.05) correlations were found between flowering and the conditionsprevailing in April to May—the main flowering season—ofthe current year. Copyright 2001 Annals of Botany Company Climate, flowering, rainfall, temperature, Anemone nemorosa, Cardamine bulbifera, Lamiastrum galeobdolon, Oxalis acetosella, Ranunculus ficaria, Stellaria holostea, Viola reichenbachiana, Viola riviniana     

家蚕保存系统红色卵的遗传分析

用遗传背景清楚的家蚕Bombyx mori红卵（re）、白卵(w-2、pe)、第4褐卵（b-4）的标志基因系统和正常型黑卵系统与我国家蚕基因库保存的20个红色卵系统杂交,进行顺反测验,分析了它们的卵色支配基因及遗传规律。结果发现：①在03-310系统中存在家蚕卵色新突变pink egg,与红卵re 等位,基因符号为re^p,表型特征为：卵淡红色,成虫蛾眼也为淡红色;②6个系统为红卵（re）的纯合系统,还有５个系统除具有re／re基因型外,还具有支配白色卵或浅红色或橙红色卵的突变基因;③２个系统为第4褐卵（b4）的纯合系统; ④6个系统的红褐色卵为母性影响遗传;⑤发现家蚕卵色基因b-4和r-e的互补关系,b-4/b-4 re/re基因型表现为新的卵色——橙黄色。     

The Scaling of Plant Height: A Comparison Among Major Plant Clades and Anatomical Grades

The scaling plant height h (m) with respect to stem diameterd (m) was determined for a total 610 species (mosses, n = 40;pteridophytes, n = 16; dicotyledonous herbs, n = 117; palms,n = 17; gymnosperms, n = 105; dicotyledonous trees, n = 315);axial length or mass vs. d was determined for the pteridophytePsilotum nudum ; and the scaling of critical buckling heighth_crit of gymnosperm and dicotyledonous trees was calculatedbased on the record trunk d and average Young's modulus E anddensity p of 33 wood species. The scaling exponents (based onleast squares and reduced major axis regressions;